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作物学报 ›› 2026, Vol. 52 ›› Issue (6): 1891-1901.doi: 10.3724/SP.J.1006.2026.53072

• 耕作栽培·生理生化 • 上一篇    下一篇

遮阴改变源-库平衡和调节碳水化合物代谢进而抑制夏玉米幼穗发育

刘恩波(), 陈静, 李红星, 于宁宁, 任佰朝, 赵斌, 刘鹏, 张吉旺*()   

  1. 山东农业大学农学院, 山东泰安 271018
  • 收稿日期:2025-09-22 接受日期:2026-02-27 出版日期:2026-06-12 网络出版日期:2026-03-06
  • 通讯作者: * 张吉旺, E-mail: jwzhang@sdau.edu.cn
  • 作者简介:刘恩波, E-mail: liuenbo5533@163.com
  • 基金资助:
    财政部和农业农村部国家现代农业产业技术体系建设专项(CARS-02-21)

Shading inhibits ear and tassel development of summer maize by altering source-sink balance and regulating carbohydrate metabolism

Liu En-Bo(), Chen Jing, Li Hong-Xing, Yu Ning-Ning, Ren Bai-Zhao, Zhao Bin, Liu Peng, Zhang Ji-Wang*()   

  1. Agronomy College of Shandong Agricultural University, Tai’an 271018, Shandong, China
  • Received:2025-09-22 Accepted:2026-02-27 Published:2026-06-12 Published online:2026-03-06
  • Contact: * Zhang Ji-Wang, E-mail: jwzhang@sdau.edu.cn
  • Supported by:
    China Agriculture Research System of MOF and MARA(CARS-02-21)

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摘要：

遮阴是影响玉米生长发育和产量形成的重要环境胁迫之一, 尤其在密植栽培或间作系统中, 玉米穗部受光照不足可能易导致显著减产。为明确遮阴对玉米幼穗发育的影响, 本研究于2023—2024年夏季在玉米穗期(拔节期至抽雄期)设置遮阴处理(S)和自然光照处理(CK), 系统测量穗部形态、穗部碳水化合物含量、激素水平及叶片的光合性能。结果显示, 遮阴显著降低夏玉米叶片的光合能力, 其中遮阴处理叶片在大喇叭口期(V12)、第15片完全展开叶期(V15)及抽雄期(VT)的蔗糖含量较CK分别下降65.0%、24.2%、23.0%, 并引起植株生物量相应降低39.5%、32.3%和41.9%。与对照(CK)相比, 遮阴导致叶片光合性能减弱, 碳水化合物合成能力下降, 进一步影响其在穗部的供应与利用, 表现为雌穗和雄穗中果糖、葡萄糖、蔗糖和淀粉含量显著降低。此外, 遮阴还显著改变发育中雌雄穗的脱落酸(ABA)、生长素(IAA)和细胞分裂素(CTK)含量, 最终导致穗发育进程延缓, 穗长和直径减少, 可育小花数量明显下降。总之, 穗期遮阴通过削弱玉米植株的同化物积累与幼穗碳水化合物代谢, 显著抑制幼穗的正常发育和形态建成, 致使穗粒数减少32.2%, 单株产量下降32.8%。本研究从生理机制层面揭示了遮阴胁迫影响玉米产量形成的过程, 为耐阴型玉米品种选育与栽培调控途径的优化提供了理论依据。

关键词: 玉米, 遮阴, 光合作用, 穗发育, 代谢

Abstract:

Shading is a major environmental stressor that strongly affects maize growth, development, and yield formation. It is particularly problematic in high-density planting and intercropping systems, where inadequate light interception around the ear can cause substantial yield losses. To examine how shading influences young ear development, field experiments were conducted during the ear development period (jointing stage to tasseling stage) in the summers of 2023 and 2024, with a shading treatment (S) and a natural-light control (CK). Ear morphology, carbohydrate contents in ears, hormone levels, and leaf photosynthetic performance were measured. Shading significantly reduced leaf photosynthetic capacity in summer maize. Relative to CK, leaf sucrose content under shading declined by 65.0%, 24.2%, and 23.0% at V12 (12th leaf stage), V15 (15th leaf stage), and VT (tasseling stage), respectively, accompanied by reductions in plant biomass of 39.5%, 32.3%, and 41.9%. The weakened photosynthetic performance and reduced assimilate production under shading further constrained carbohydrate supply to, and utilization within, developing ears, resulting in significantly lower fructose, glucose, sucrose, and starch contents in both female and male ears. Shading also markedly altered abscisic acid (ABA), auxin (IAA), and cytokinin (CTK) levels in developing ears, leading to delayed ear development, smaller ear length and diameter, and a pronounced reduction in fertile floret number. Overall, shading during the ear stage inhibited normal ear development by limiting assimilate accumulation and disrupting carbohydrate metabolism in young ears, which contributed to a 32.2% reduction in kernels per ear and a 32.8% decrease in yield per plant. These findings clarify the physiological basis of shading-induced yield loss in maize and provide a foundation for breeding shade-tolerant cultivars and optimizing cultivation practices.

Key words: corn, shading, photosynthesis, ear development, metabolism

表1

遮阴对田间小气候的影响"

年份
Year		 处理
Treatment		 光照强度
Light intensity (μmol m-2 s-1)		 风速
Air speed
(m s-1)		 温度
Temperature
(℃)		 相对湿度
Relative humidity
(%)		 CO2浓度
CO2 concentration
(μmol mol-1)
顶部
Top		 穗部
Ear		 底部
Bottom
2023 S 463.73 b 165.70 b 102.63 a 0.14 a 30.23 a 72.58 a 308.09 a
CK 1142.40 a 300.70 a 107.53 a 0.08 a 31.21 a 73.67 a 303.17 a
2024 S 428.04 b 134.81 b 84.10 a 0.20 a 31.87 a 88.72 a 326.21 a
CK 920.85 a 246.26 a 85.30 a 0.11 a 32.60 a 87.37 a 328.78 a

表2

遮阴对夏玉米收获时穗部和产量相关性状的影响"

年份
Year		 处理
Treatment		 穗粒数
Grains per ear		 千粒重
1000-grain weight (g)		 籽粒产量
Grain yield (g plant-1)		 果穗长度
Ear length (cm)		 果穗直径
Ear diameter (cm)
2023 S 376.54±4.09* 371.72±1.41 139.97±1.16* 15.74±0.63* 4.31±0.23*
CK 561.28±12.19 374.76±3.22 209.76±1.85 21.04±0.28 5.23±0.27
2024 S 384.19±7.31* 340.26±12.19 130.73±3.20* 17.51±0.24* 4.35±0.04*
CK 561.37±2.36 343.45±1.95 192.80±0.45 20.71±0.38 4.71±0.06

表3

遮阴对不同生育时期生物量积累及其所占比例的影响"

年份
Year		 生育
时期
Growth stage		 处理
Treatment		 叶片Leaf 雌穗Ear 雄穗Tassel 总干物质量
Total dry
mass
(g plant-1)
干物质量
Dry mass
(g plant-1)		 占比Proportion
(%)		 干物质量
Dry mass
(g plant-1)		 占比Proportion
(%)		 干物质量
Dry mass
(g plant-1)		 占比Proportion
(%)
2023 V12 S 21.08±3.81* 57.62 0.01±0.00* 0.02 0.40±0.07* 1.12 36.38±4.13*
CK 31.82±0.69 52.89 0.03±0.01 0.05 1.03±0.83 1.72 60.20±1.91
V15 S 25.55±1.52* 41.94 0.17±0.04* 0.28 1.05±0.04* 2.24 60.61±2.96*
CK 36.96±1.11 41.54 0.39±0.16 0.44 3.79±0.78 4.26 89.13±5.16
VT S 25.90±0.86* 40.53 0.76±0.07* 1.19 1.36±0.08* 1.64 64.23±1.41*
CK 42.30±1.41 39.11 1.31±0.14 1.25 4.33±1.04 3.95 107.21±2.67
2024 V12 S 18.82±1.02* 59.14 0.01±0.00* 0.03 0.44±0.05* 1.39 31.81±1.45*
CK 27.10±1.98 51.62 0.05±0.02 0.10 1.08±0.11 2.06 52.54±3.71
V15 S 21.85±1.47* 45.28 0.21±0.02* 0.43 0.80±0.24* 1.66 48.33±4.05*
CK 31.26±3.10 42.71 0.38±0.03 0.52 2.36±0.39 3.24 73.14±6.06
VT S 24.23±0.92* 41.56 0.80±0.08* 1.37 1.11±0.13* 1.91 58.30±0.95*
CK 40.72±0.93 39.32 1.47±0.12 1.42 4.15±0.15 4.00 103.63±2.83

图1

遮阴对夏玉米雌穗和雄穗不同时期发育情况的影响 A和B分别是雄穗、雌穗形态对比。处理同表1。V9: 小喇叭口期; V10: 第10片展开叶期;V12: 大喇叭口期; V15: 第15片完全展开叶期; VT: 抽雄期。"

图2

遮阴对夏玉米雌穗和雄穗大小的影响 处理同表1。缩写同表3。不同小写字母表示不同处理间差异显著(P < 0.05)。"

表4

遮阴对雄穗小花发育特性的影响"

年份
Year		 处理
Treatment		 总小花数
Number of tassel flowers (per plant)		 不育小花数
Aborted tassel flowers
(per plant)		 分支数
Number of tassel branches (per plant)		 不育率
Abortion rate
(%)
2023 S 451.33±42.96* 156.33±10.98 5.55±1.09* 34.88±3.91
CK 831.33±23.44 4.67±2.94* 9.16±1.16 0.56±0.35*
2024 S 446.17±43.41* 148.33±19.61 6.17±0.75* 33.67±6.64
CK 743.00±29.44 5.50±1.52* 9.00±1.09 0.74±0.20*

表5

遮阴对雌穗小花发育特性的影响"

年份
Year		 处理
Treatment		 总小花数
Total floret number
(per ear)		 受精小花数
Fertilized florets
(per ear)		 穗粒数
Kernels
(per ear)		 小花受精率
Floret
fertilization rate (%)		 小花结实率
Floret Setting rate (%)		 籽粒败育率
Grain abortion
rate (%)		 总结实率Total setting
rate (%)
2023 S 653.33±29.36* 519.33±12.83* 376.54±4.09* 79.61±3.73* 72.54±1.77 27.46±1.77 57.07±1.75
CK 975.00±21.04 822.17±28.18 561.28±12.19 84.32±1.73 68.33±2.33 31.67±2.33 56.82±1.22
2024 S 593.00±48.72* 473.83±16.06* 384.19±7.31* 78.24±3.99* 81.16±2.77 18.84±2.77 65.10±5.29
CK 912.00±47.85 750.00±41.57 561.37±2.36 82.24±1.63 75.05±4.30 24.95±4.30 62.59±2.86

图3

遮阴对不同生育时期叶片光合性能相关参数的影响 处理同表1。缩写同表3。不同字母表示同一时期S与CK差异显著(P < 0.05)。"

图4

遮阴对不同生育时期叶片蔗糖合成酶(SS)活性、蔗糖磷酸合成酶(SPS)活性和蔗糖含量的影响 处理同表1。缩写同表3。不同字母表示同一时期S与CK差异显著(P < 0.05)。"

图5

遮阴对不同生育时期穗部单位鲜重碳水化合物含量的影响 处理同表1。缩写同表3。不同字母表示同一时期S与CK差异显著(P < 0.05)。"

图6

遮阴对不同生育时期雌穗和雄穗中细胞壁转化酶(CWI)和可溶性淀粉合成酶(SSS)活性的影响 处理同表1。缩写同表3。不同字母表示同一时期S与CK差异显著(P < 0.05)。"

图7

遮阴对不同时期雌穗和雄穗中脱落酸(ABA)、生长素(IAA)、细胞分裂素(CTK)含量的影响 处理同表1。缩写同表3。不同字母表示同一时期S与CK差异显著(P < 0.05)。"

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